A) The bacterium is spatially discretized into 0.1 µm long mesh elements along the long-axis (the x-direction). In each of the mesh elements, two state variables and –the number of barbed-ends and the f-actin density, respectively– determine the element contribution to propulsion force and drag coefficient. B) Equations for , for example, are derived from conservation of barbed ends at a single mesh element: , where is the nucleation of new barbed-ends (i.e. a new mother filament), is the autocatalytic creation of barbed-ends from existing barbed-ends (i.e. branching), and is the velocity of the bacterium. Only elements on the hemispherical cap significantly contribute to propulsion force, while f-actin along the side of the bacterium contributes to the drag force. The ratio of force to drag determines the instantaneous velocity . A similar element diagram can be drawn for , though without any autocatalytic term. C) Steady state speeds for different ActA distributions. A constant drag on the bacterium led to faster normals than ultrapolars. Incorporation of a linear filament-dependent restraining mechanism, representing either (or both) filament-surface tethers or fluid coupling led to faster ultrapolars than normals, as in our experimental observations. A cooperative filament-dependent restraining mechanism (representing kinetic friction, for example) similarly led to faster ultrapolars (data not shown).